The influence of magnesium deficiency on carbohydrate concentrations in Norway spruce (Picea abies) needles

Both short- and long-term effects of Mg deficiency on carbohydrate metabolism were investigated in 6-year-old clonal Norway spruce (Picea abies (L.) Karst.) trees cultivated in sand culture with an optimal supply of nutrients, except for Mg which was supplied at 0.203, 0.041 and 0.005 mM to provide optimal, moderately deficient and severely deficient Mg supply, respectively. Annual changes in carbohydrate concentrations (starch, sucrose, glucose and fructose) were analyzed and diurnal changes were investigated on a single day during the summer. Older needles of trees in the moderate Mg-deficiency treatment developed tip-yellowing symptoms, whereas current-year needles remained green. The severe Mg-deficiency treatment led to pronounced yellowing symptoms in needles of all ages. Increased carbohydrate concentrations were observed before needle yellowing occurred. Diurnal and annual changes in carbohydrates were similar in all treatments; however, carbohydrate concentrations were influenced by Mg supply. In both Mg-deficiency treatments, starch concentrations increased in needles, especially during summer and autumn. Starch accumulation was more pronounced at the beginning of the Mg-deficiency treatments than at the end of the treatments. Sucrose, and to a minor extent, glucose and fructose concentrations tended to increase in response to Mg deficiency. The consequences of Mg deficiency on carbohydrate metabolism are discussed with respect to reduced plant growth and decreased transport rates of carbohydrates to sink organs.     

Uptake and distribution of calcium and phosphorus in beech (Fagus sylvatica) as influenced by aluminum and nitrogen

We studied the effects of excess nitrogen added as nitrate (NO(3) (-)) or ammonium (NH(4) (+)), or both, on mineral nutrition and growth of beech (Fagus sylvatica L.) plants grown at pH 4.2 in Al-free nutrient solution or in solutions containing 0.1 or 1.0 mM AlCl(3). A high external concentration of NH(4) (+) increased the concentration of nitrogen in roots, stems and leaves. The root/shoot dry weight ratio was less in plants grown in the presence of NH(4) (+) than in plants grown in the presence of NO(3) (-). The concentration of phosphorus in the roots was increased and the concentration of potassium in all parts of the plant was decreased by NH(4) (+). A high external concentration of NO(3) (-) caused a decrease in phosphorus concentrations of the root, stem and leaf. Uptake of (45)Ca(2+) by roots was reduced in the presence of high concentrations of NH(4) (+) or NO(3) (-), and a combination of high concentrations of nitrogen and aluminum further reduced the uptake of (45)Ca(2+). Uptake of phosphate ((32)P) and concentrations of phosphorus in root and shoot were increased when plants were grown in the presence of 0.1 mM Al. Exposure to 1.0 mM Al, however, reduced the concentration of phosphorus in roots and shoots and the reduction was greater when plants were grown in the presence of a high external NO(3) (-) concentration. Aluminum binds to roots, and plants grown in the presence of 1.0 mM Al had a slightly higher concentration of aluminum in roots than plants grown in the presence of 0.1 mM Al, whereas the concentration of Al in the shoot was increased 2 to 3 times in plants exposed to 1.0 mm Al. Furthermore, the effects of 1.0 mM Al on uptake of other macronutrients were quite different from the effects of 0.1 mM Al. We conclude that 0.1 mM Al facilitates uptake and transport of phosphorus in beech and that between 0.1 and 1.0 mM Al there is a dramatic change in the effects of Al on uptake and transport of divalent cations and phosphorus.     

Physiological responses of black spruce layers and planted seedlings to nutrient addition

We investigated effects of nutrient addition on several physiological characteristics of 60-cm-tall black spruce (Picea mariana Mill. B.S.P.) layers (i.e., rooted branches of overstory trees) and 20-cm-tall planted seedlings on a clear-cut, N-limited boreal site. After two growing seasons, current-year and one-year-old needles of fertilized trees (layers and seedlings combined) had higher net photosynthetic rates (A(n)) and maximum capacity of Rubisco for CO(2) fixation (V(max)) than unfertilized trees. One-year-old needles of fertilized trees had higher stomatal conductance (g(s)), higher water-use efficiency, and lower intercellular to ambient CO(2) ratio than unfertilized trees. Additionally, fertilized trees had higher predawn and midday shoot water potentials than unfertilized trees. Stomatal conductance of 1-year-old needles was 23% higher in seedlings than in layers, but there were no significant differences in g(s) of current-year needles between the regeneration types. For both needle age-classes, A(n) and V(max) of layers were 25 and 40% higher, respectively, than the corresponding values for seedlings. The higher values of A(n), V(max) and foliar N concentration of layers compared with seedlings after two growing seasons may be associated with the larger root systems of the layers compared with the transplanted seedlings.     

Effect of mineral nutrition content on oxygen exchange and chlorophyll a fluorescence in needles of Norway spruce

Photosynthetic O(2) evolution at high irradiances (approximately 600-1000 micro mol m(-2) s(-1)) and O(2) uptake in darkness were measured in needles of control, irrigated and irrigated-fertilized trees of Norway spruce (Picea abies (L.) Karst.). Measurements were made at 20 degrees C and at high CO(2) concentrations. The results suggest that, at given times of the year, a major part of the variation in gross photosynthesis of current-year and one-year-old needles across treatments is associated with differences in needle N content. Furthermore, the rate of O(2) uptake measured after 5 or 10 min in darkness was positively correlated with both the preceding rate of gross O(2) evolution and the N content in fully expanded current-year needles. Measurements of chlorophyll a fluorescence, taken simultaneously with measurements of O(2) evolution in current-year sun needles, showed that Stern-Volmer quenching of minimum fluorescence and the ratio of variable to maximum fluorescence in the dark- and light-adapted state were strongly correlated with the gross rate of O(2) evolution. This suggests that the increased rate of gross photosynthesis in needles of irrigated-fertilized trees was associated with adjustments in the thermal energy dissipation within photosystem II.     

Storage and internal cycling of nitrogen in relation to seasonal growth of Sitka spruce

Three-year-old clonal cuttings of Picea sitchensis (Bong.) Carr. were grown for two years (1988-1989) in sand irrigated with a nutrient solution containing either 1.0 mol N m(-3) (low N) or 6.0 mol N m(-3) (high N) NH(4)NO(3). In 1988, all the N provided was enriched with (15)N to 4.95 atom % (labeled N). In 1989, N was supplied with (15)N at natural abundance (unlabeled N). The recovery of unlabeled and labeled N in new foliage was used to quantify the internal cycling of N. In the high-N treatment, trees had two flushes of shoot growth and a period of rapid root growth, which coincided with the second flush of shoot growth in August. The timing of root growth and the first flush of shoot growth was similar in the low-N treatment, but there was no second flush of shoot growth and a greater proportion of biomass was recovered in roots. By November 1989, the root/needle dry matter ratio was 1.95 for the low-N trees and 1.36 for the high-N trees. Nitrogen was stored overwinter in roots and current-year needles. During the first six weeks of growth in the spring of 1989, stored N was remobilized for new foliage growth. Subsequent growth depended on root uptake of N. Remobilization of stored N was apparently not affected by the current N supply, because the amount of unlabeled N recovered in foliage produced in 1988 was the same for both N treatments. During 1989, the proportion of (15)N remobilized from roots relative to that from leaves produced in 1988 was greater in low-N trees than in high-N trees. In the autumn of both years, there was rapid uptake of N into roots and current-year needles. The effects of N supply on tree growth and nitrogen use efficiency are discussed in terms of the capacity for both N storage and internal cycling.     

Effects of cone-induction treatments on black spruce (Picea mariana) current-year needle development and gas exchange properties

Both drought and root pruning (RP) increased the number of cones induced when black spruce (Picea mariana (Mill.) B.S.P.) grafts were injected with gibberellins A(4/7) (GA), but their effects on predawn shoot water potential and current-year needle development differed. Drought decreased predawn shoot water potential (Psi(pd)), but only during the period when irrigation was withheld, and it had no effect on the growth or gas exchange properties of current-year needles. Conversely, root pruning had little effect on Psi(pd), but it resulted in trees with smaller current-year needles that had lower nitrogen and chlorophyll concentrations and reduced rates of gas exchange up to the later stages of shoot elongation compared with needles of control trees. These findings are discussed in relation to potential effects on the development of induced cones in the following growth cycle.     

Photosynthesis,nutrient, growth and soil investigations of a declining Norway spruce (Picea abies) stand in the coastal region of Northern Germany

The research site, Wingst Compartment 123B, is a 68-year-old Norway spruce (Picea abies (Karst.)) stand located in the coastal area of northern Germany. This area receives high atmospheric inputs of ammonium and also has relatively high ozone concentrations (0.061 mg m⁻³).Ten trees were categorized as healthy to slightly damaged (3–29% needle loss) or severely damaged (49–71% needle loss). Apparent net photosynthetic rates were measured on detached branches at light saturation (1000 μE m⁻² s⁻¹). Needles were analyzed for chlorophyll, N, C, Ca, Mg, K, Zn, Mn and Fe.When compared to the healthy-to-slightly-damaged trees, the severely damaged trees tended to have higher rates of net apparent photosynthesis in the 1 and 2-year-old needles and similar rates in the current-year needles. All three needle ages from the severely damaged trees had higher average stomatal conductances to water vapor (g_s. Although the damaged trees had significantly less total chlorophyll in all needle ages sampled, there was no statistically significant difference in the chlorophyll a:b ratio between the healthy and severely damaged trees.Nitrogen contents of the current-year needles were slightly lower in the severely damaged trees. Carbon and calcium levels did not significantly differ between the damage classes, although the average Ca content of all younger needles was generally under the recommended sufficiency levels. The current-year needles of the severely damaged trees had significantly less magnesium (0.42–0.46 mg Mg g⁻¹) than those of the healthy trees, and all trees had Mg values in the deficiency range (< 0.7 mg g⁻¹). The severely damaged trees also had lower average potassium levels in the older needles.The annual volume increment per unit crown surface area declined with increasing crown damage. Trees with a 50% needle loss showed a 62% loss of volume increment.Soil investigations revealed conditions of high soil acidity and poor nutrient capacity. The low pH values (pH < 3.8) in 64% of all samples indicated a high risk of acid toxicity for plant roots in the investigated area.The significance of these results relative to the current ideas concerning forest decline is discussed.     

Nitrate content, amino acid composition and growth of yellow birch seedlings in response to light and nitrogen source

Yellow birch (Betula alleghaniensis Britt.) seedlings were grown for three months in a greenhouse at two radiant flux densities-full light (FL) and 50% shade (LL)-and with three nitrogen sources- ammonium only (NH(4) (+)), nitrate only (NO(3) (-)) and a 1:1 mixture of ammonium and nitrate (NH(4) (+)/NO(3) (-))-in a completely randomized factorial design. The total biomass of seedlings grown under low light (LL) did not vary significantly with nitrogen source; although NO(3) (-)-treated seedlings were smaller and had a significantly lower (P 相似文献   

Seasonal nutrient dynamics in white pine and white spruce in response to environmental manipulation

Seasonal retranslocation in white pine (Pinus strobus L.) and white spruce (Picea glauca (Moench) Voss) was examined in response to silvicultural treatments (scarification, annual fertilization application, and annual control of competing vegetation with herbicide) that changed both environmental conditions and the growth rate of the trees. Four years after plantation establishment and initial treatment, nutrient accumulation in current-year needles of white pine and retranslocation from 1-year-old needles were increased following the vegetation control treatment, which increased resource availability (nutrients, water and light) and, hence, growth rate. Nutrient accumulation also increased in current-year white spruce needles following the same treatment, whereas retranslocation decreased in 1-year-old white spruce needles. Correlations of retranslocation (N, P and K) with growth rate (shoot biomass increment) showed a strong positive relationship for white pine and a negative relationship for white spruce. Retranslocation of K was correlated with foliar and soil K concentrations; the availability of this nutrient was also significantly reduced by vegetation control. A general theory for the control of nutrient retranslocation in conifers, which is not based exclusively on either sink strength or soil nutrient availability, is proposed. We conclude that retranslocation response is species specific and related to the potential phenotypic growth response to changing environmental conditions and to short-term imbalance in the supply versus the demand for nutrients.     

Internal cycling of nitrogen, potassium and magnesium in young Sitka spruce

Potassium (K) and magnesium (Mg) are essential macro-nutrients, but little is known about how they are cycled within plants. Stable isotope studies have shown that the internal cycling of nitrogen (N) is independent of current nutrient supply in temperate tree species. This is ecologically significant because it allows trees to produce rapid shoot growth in spring independent of current soil N uptake. We used stable isotopes to quantify N, K and Mg in new shoots of Sitka spruce (Picea sitchensis (Bong.) Carr.) seedlings and to compare the relative contributions from current uptake and internal cycling. Two-year-old Sitka spruce seedlings were labeled with (15)N, (41)K and (26)Mg in an abundant or a limited supply for one growing season. The trees were repotted in the subsequent dormant season to prevent further root uptake of enriched isotopes and provided with an abundant or a limited supply of unlabeled nutrients until they were harvested in early summer of the following year. The supply was switched for half the trees in the second year to create four nutrient regimes. Enrichment of (15)N, (41)K and (26)Mg in current-year growth was attributed to internally cycled N, K and Mg uptake from the previous year. The internal cycling of N, K and Mg in new growth was significantly affected by the first-year nutrient treatments. The second-year nutrient supply affected the growth rates of the trees, but had no effect on the amounts of N, K or Mg contributed from internal cycling. Thus, internal cycling of K and Mg in Sitka spruce are, like that of N, independent of current nutrient supply.     

Influence of a high Mn supply on Norway spruce (Picea abies (L.) Karst.) seedlings in relation to the nitrogen source

Effects of 3, 25, 100, 200 and 800 microM Mn on biomass and pigment, starch and nitrate concentrations were studied in Norway spruce (Picea abies (L.) Karst.) seedlings grown with either NO(3) (-) or NH(4) (+) as the sole nitrogen source. After 77 days of exposure to 800 microM Mn, shoot growth had ceased in about 50% of the seedlings independently of the N source. Despite high Mn concentrations in roots and shoots of the Mn-treated seedlings, no visible symptoms of Mn toxicity were evident. The rate of root elongation was decreased by treatment with >/= 200 microM Mn when N was supplied as NO(3) (-), but not when it was supplied as NH(4) (+). This difference could be attributed to the higher Mn concentrations in root tips of the NO(3) (-)-grown seedlings compared with the NH(4) (+)-grown seedlings. In Mn-treated seedlings, the concentration of Mg, and to a lesser extent that of Ca, decreased. Depletion of these elements might account for the observed growth depression. Potassium concentrations were similar in the control and Mn-treated seedlings. Treatment of seedlings with 800 microM Mn for 50 days led to several physiological changes: starch accumulated, the concentrations of nitrate and phenolic compounds increased, pigment concentrations decreased, and in vivo nitrate reductase activity in roots was reduced.     

Effects of elevated CO(2) concentration and nutrition on net photosynthesis,stomatal conductance and needle respiration of field-grown Norway spruce trees

To study the effects of elevated CO(2) on gas exchange, nonstructural carbohydrate and nutrient concentrations in current-year foliage of 30-year-old Norway spruce (Picea abies (L.) Karst.) trees, branches were enclosed in ventilated, transparent plastic bags and flushed with ambient air (mean 370 &mgr;mol CO(2) mol(-1); control) or ambient air + 340 &mgr;mol CO(2) mol(-1) (elevated CO(2)) during two growing seasons. One branch bag was installed on each of 24 selected trees from control and fertilized plots. To reduce the effect of variation among trees, results from each treated branch were compared with those from a control branch on the same whorl of the same tree. Elevated CO(2) increased rates of light-saturated photosynthesis on average by 55% when measured at the treatment CO(2) concentration. The increase was larger in shoots with high needle nitrogen concentrations than in shoots with low needle nitrogen concentrations. However, shoots grown in elevated CO(2) showed a decrease in photosynthetic capacity compared with shoots grown in ambient CO(2). When measured at the internal CO(2) concentration of 200 &mgr;mol CO(2) mol(-1), photosynthetic rates of branches in the elevated CO(2) treatments were reduced by 8 to 32%. The elevated CO(2) treatment caused a 9 to 20% reduction in carboxylation efficiency and an 18% increase in respiration rates. In response to elevated CO(2), starch, fructose and glucose concentrations in the needles increased on average 33%, whereas concentrations of potassium, nitrogen, phosphorus, magnesium and boron decreased. Needle nitrogen concentrations explained 50-60% of the variation in photosynthesis and CO(2) acclimation was greater at low nitrogen concentrations than at high nitrogen concentrations. We conclude that the enhanced photosynthetic rates found in shoots exposed to elevated CO(2) increased carbohydrate concentrations, which may have a negative feedback on the photosynthetic apparatus and stimulate cyanide-resistant respiration. We also infer that the decrease in nutrient concentrations of needles exposed to elevated CO(2) was the result of retranslocation of nutrients to other parts of the branch or tree.     

Growth, allocation and tissue chemistry of Picea abies seedlings affected by nutrient supply during the second growing season

One-year-old Norway spruce (Picea abies (L.) Karst.) seedlings were grown hydroponically in a growth chamber to investigate the effects of low and high nutrient availability (LN; 0.25 mM N and HN; 2.50 mM N) on growth, biomass allocation and chemical composition of needles, stem and roots during the second growing season. Climatic conditions in the growth chamber simulated the mean growing season from May to early October in Flakaliden, northern Sweden. In the latter half of the growing season, biomass allocation changed in response to nutrient availability: increased root growth and decreased shoot growth led to higher root/shoot ratios in LN seedlings than in HN seedlings. At high nutrient availability, total biomass, especially stem biomass, increased, as did total nonstructural carbohydrate and nitrogen contents per seedling. Responses of stem chemistry to nutrient addition differed from those of adult trees of the same provenance. In HN seedlings, concentrations of alpha-cellulose, hemicellulose and lignin decreased in the secondary xylem. Our results illustrate the significance of retranslocation of stored nutrients to support new growth early in the season when root growth and nutrient uptake are still low. We conclude that nutrient availability alters allocation patterns, thereby influencing the success of 2-year-old Norway spruce seedlings at forest planting sites.     

氮肥对盆栽欧洲水青冈生物量和营养分配的影响(英文)

调查了施加氮肥(15NH4和15NO3)处理后在两个连续生长季内欧洲水青冈(Fagus sylvatica L.)幼苗地上部分和地下部分的生物量和营养元素分配。盆栽欧洲水青冈幼苗培养于温室大棚内,培养土样取自相邻的三种林分:欧洲水青冈,挪威赤松,欧洲水青冈-赤松混交林。结果表明,氮肥(15N)处理对欧洲水青冈营养元素分配没有显著影响,施加氮素形式决定自身流入植物库的情况。欧洲水青獭收氮素主要以硝态氮的形式,因此,尽管植物体内保存的硝态氮和氨态氮并没有统计差异,但是叶片中硝态氮明显减少。施加硝态氮对欧洲水青冈氮素恢复的影响要大于施加氨态氮。与欧洲水青冈茎、粗根相比,优质根系对氮素(15N)固定是一个缓慢过程。表8图1参40。     

Effects of chronic acid fog exposure with soil acidification or nitrogen loading on nutrient status in Abies firma seedlings

To clarify the nutrient status in momi fir (Abies firma Sieb. et Zucc.) seedlings under complex stress conditions of acid fog with soil acidification or nitrogen loading, we exposed seedling shoots to simulated acid fog (pH 3) and simultaneously loaded the soil with acid or excess nitrogen for 17 months. Proton and nitrogen loading reduced K concentration in soil, but these treatments had little effect on the nutrient status of fine roots in momi fir seedlings. Acid fog exposure resulted in reduced concentrations of Mg, Al, and Fe in current-year needles and Mg, Al, Cu, Fe, and Zn in 1-year-old needles. The complex effects of acid fog exposure with proton or nitrogen loading on nutrient status were relatively slight. However, elements such as Al and Cu notably reflect the effects of proton loading. These results show that the nutrient status of trace elements, rather than major elements, in needles of momi fir seedlings is sensitive to exposure to acid fog or proton loading.     

Absorption and assimilation of nitrate and ammonium ions by jack pine seedlings

Jack pine (Pinus banksiana Lamb.) seedlings were grown in a shaded or unshaded light regime with either NO(3) (-)- or NH(4) (+)-N as the sole N source. After three months, seedlings grown with NH(4) (+)-N were larger than seedlings grown with NO(3) (-)-N. Irradiance had a greater effect on growth of ammonium-fed seedlings than on growth of nitrate-fed seedlings.At all times from 6 to 24 h following incorporation of (15)N, soluble, insoluble, and total (15)N contents of shoots and roots were higher in ammonium-fed seedlings than in nitrate-fed seedlings. The pattern of (15)N accumulation in shoots was similar to that in roots. After 6 and 24 h of (15)N incorporation, unshaded, ammonium-fed seedlings had 8.8 and 2.8 times greater total (15)N contents, respectively, than unshaded, nitrate-fed seedlings. In response to shading, ammonium-fed seedlings increased their total uptake of (15)N per unit root weight, whereas nitrate-fed seedlings did not. No nitrate or (15)NO(3) (-) was detected in any plant tissue. Nitrate-fed plants had higher NH(4) (+), Asp, and Gln concentrations in needles and higher gamma-aminobutyric acid and Arg concentrations in stems. Accumulation of (15)N in roots was not affected by the pH of the (15)N solution or by the N source fed to the seedlings before the period of (15)N incorporation. Thus NO(3) (-) transport into roots, rather than its reduction or transport within the plant, seems to be the factor limiting the growth of jack pine supplied with NO(3) (-)-N as the sole N source.     

Defoliation and growth relationships for mid-rotation Sitka spruce attacked by the green spruce aphid, Elatobium abietinum (Walker) (Homoptera: Aphididae)

Green spruce aphid (Elatobium abietinum) feeds primarily on the 1-year-old and older needle leaves of spruce (Picea spp.) and is a major defoliator of commercial Sitka spruce plantations in the UK and other maritime regions of north-west Europe. The impact of E. abietinum on mid-rotation (23-28-year-old) Sitka spruce in Radnor Forest, in mid-Wales, was determined by comparing aphid population densities, defoliation and growth between plots of trees that were either treated with insecticide or were left untreated. The experimental treatments were maintained for 5 years and over this period (2002-2006) peak E. abietinum densities in the untreated plots varied from 5 to 36 aphids per 100 needles. These densities, which were representative of low to moderate rates of infestation, were associated with low rates of defoliation (0-8%), but they had a significant (P ? 0.01) impact on mean radial increment (RI) and mean volume increment (VI). In 2005, the year with the highest aphid populations, peak densities in the untreated plots averaged 14 aphids per 100 needles and this rate of infestation reduced RI by 17%, VI by 10% and the dry weight of current-year needles by 10%. On average, across all years, infestation by E. abietinum reduced mean annual VI by 6%. Comparisons with previous studies on the impact of the aphid on 4-year-old, 7-year-old and 9-year-old Sitka spruce (Straw et al., 2005) indicate that defoliation by E. abietinum, on a per capita basis, decreases as trees grow older, but that the impact on VI increases. The different growth response of young and mid-rotation Sitka spruce to infestation is related to differences in canopy structure, particularly in the ratio of current-year needles to older needles, and the greater demands on photosynthetic production in older trees that arise from the need to support an increasing quantity of non-photosynthetic structural tissues in branches, stem and roots.     

Rapid freezing injury in red spruce: seasonal changes in sensitivity and effects of temperature range

On calm, cold days in winter, sun-exposed needles of red spruce (Picea rubens Sarg.) may warm 10 to 20 degrees C above ambient air temperature, and undergo rapid (>/= 1 degrees C min(-1)) fluctuations in temperature as light breezes or passing clouds alter the energy balance of the foliage. It has been proposed that the resulting rapid freeze-thaw cycles (freezing stress) cause a type of winter injury in montane red spruce that is characterized by necrosis of sun-exposed foliage. In autumn and winter, we monitored rapid freezing stress response of needle sections from 10 montane red spruce trees by subjecting needles to rapid freezing over the temperature span typically recorded in the field. In autumn, experimental rapid freezing stress produced severe injury only at temperatures considerably lower than expected for that time of year. In winter, rapid freezing caused occasional, moderate injury in fully hardened foliage of trees susceptible to both slow and rapid freezing. Seasonal changes in sensitivity to rapid and slow freezing were correlated, suggesting that environmental factors that are known to affect sensitivity to slow freezing may also affect sensitivity to rapid freezing. Experimental manipulation of the start and end temperatures of rapid freezing stress events showed that moderate to severe needle injury can occur in susceptible trees at temperature spans slightly more extreme than those typically recorded in the field. The extent of injury was similar at different starting temperatures if rapid freezing occurred over the same temperature span. Year-old foliage was consistently less sensitive to rapid freezing stress than current-year foliage, but some year-old foliage was damaged when the rapid freezing stress regime caused severe injury in current-year foliage. We conclude that rapid freeze-thaw cycles can explain light to moderate injury of current-year foliage, but they do not explain the more severe and widespread pattern of foliar damage that has occurred intermittently over at least the last 18 years.     

Influence of nutrient supply on shade-sun acclimation of Picea abies seedlings: effects on foliar morphology,photosynthetic performance and growth

Norway spruce seedlings (Picea abies Karst.) were grown in low light for one year, under conditions of adequate and limiting nutrition, then transferred to high light. Three months after transfer we measured photosynthesis, leaf nitrogen concentration, leaf chlorophyll concentration and leaf mass per area (LMA) of current-year and 1-year-old shoots; silhouette area ratio (SAR, the ratio of shoot silhouette area to projected needle area) was also measured in current-year shoots. At the foliage level, the effects of light and nutrient treatments differed markedly. Light availability during foliage expansion primarily affected LMA and SAR (morphological acclimation at the needle and shoot level, respectively). By contrast, nutrient supply in high light affected photosynthetic capacity per unit of leaf tissue (physiological acclimation at the cellular level) but did not affect LMA and SAR. The capacity for shade-sun acclimation in foliage formed before transfer to high light differed greatly from that of foliage formed following the transfer. The morphological inflexibility of mature needles (measured by LMA) limited their shade-sun acclimation potential. In contrast, at high nutrient supply, shoots that developed just after the change in photosynthetic photon flux density largely acclimated, both morphologically and physiologically, to the new light environment. The acclimation response of both current- and 1-year-old shoots was prevented by nutrient limitation. Analysis of growth at the whole-plant level largely confirmed the conclusions drawn at the shoot level. We conclude that nutrient shortage subsequent to the opening of a canopy gap may strongly limit the acclimation response of Norway spruce seedlings. Successful acclimation was largely related to the plant's ability to produce sun foliage and adjust whole-plant biomass allocation rapidly.     

Effect of raw humus under two adult Scots pine stands on ectomycorrhization, nutritional status, nitrogen uptake, phosphorus uptake and growth of Pinus sylvestris seedlings

Ectomycorrhiza (EM) formation improves tree growth and nutrient acquisition, particularly that of nitrogen (N). Few studies have coupled the effects of naturally occurring EM morphotypes to the nutrition of host trees. To investigate this, pine seedlings were grown on raw humus substrates collected at two forest sites, R2 and R3. Ectomycorrhiza morphotypes were identified, and their respective N uptake rates from organic (2-(13)C, (15)N-glycine) and inorganic ((15)NH(4)Cl, Na(15)NO(3), (15)NH(4)NO(3), NH(4)(15)NO(3)) sources as well as their phosphate uptake rates were determined. Subsequently, the growth and nutritional status of the seedlings were analyzed. Two dominant EM morphotypes displayed significantly different mycorrhization rates in the two substrates. Rhizopogon luteolus Fr. (RL) was dominant in R2 and Suillus bovinus (Pers.) Kuntze (SB) was dominant in R3. (15)N uptake of RL EM was at all times higher than that of SB EM. Phosphate uptake rates by the EM morphotypes did not differ significantly. The number of RL EM correlated negatively and the number of SB EM correlated positively with pine growth rate. Increased arginine concentrations and critical P/N ratios in needles indicated nutrient imbalances of pine seedlings from humus R2, predominantly mycorrhizal with RL. We conclude that different N supply in raw humus under Scots pine stands can induce shifts in the EM frequency of pine seedlings, and this may lead to EM formation by fungal strains with different ability to support tree growth.